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Genetic Integrity of Long's Bulrush at Lac De L'école on the Tusket River

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Genetic Integrity of Long's Bulrush at Lac De L'école on the Tusket River Report: Genetic Integrity of Long’s bulrush at Lac de l’École on the Tusket River Nova Scotia Species at Risk Conservation Fund project number: NSSARCF09_05 Ron MacKay, Ph.D. Department of Biology Mount Saint Vincent University 166 Bedford Highway Halifax, Nova Scotia B3M 2J6 (902) 457-6293 [email protected] 31 May 2011 1 Contents Introduction………………………………………………………………………....... 2 Logistics and Methods………………………………………………………………. 7 Preliminary Results………………………………………………………………….. 10 Summary and Prospects……………………………………………………………. 13 Acknowledgements………………………………………………………………….. 16 References……………………………………………………………………………. 17 2 Introduction Long’s bulrush (Scirpus longii Fernald, Cyperaceae) is a wetland species of interest to botanists and conservationists. Members of the genus are rhizomatous perennials growing in dense mats and/or tussocks. Most species produce flowering culms on an annual basis (Whittemore and Schuyler 2003). S. longii is unique, directing more of its resources to the growth and extension of its rhizomes, producing large and long-lived clones (Hill and Johansson 1992), and flowering only when disturbed (as by fire, see Schuyler and Stasz 1985). As Long’s bulrush allocates more of its resources to vegetative growth and inhabits continuously wet and peaty soils, it has been classified (Hill and Johansson 1992) as a stress-tolerant plant (Grime 1977) adapted to low concentrations of oxygen and other nutrients. However, the induction of flowering by disturbance indicates a facultative ruderal or weedy character (Grime 1977). Thus, Long’s bulrush is adapted to an intermediate place in oligotrophic wetland succession, doing well once a peaty substratum is established, contributing to the development of a richer and drier soil, re-establishing itself (from seed and rhizome) quickly after drought and/or fire, and being excluded, eventually, by shrubs and trees. The sister species of Long’s bulrush, the wooly bulrush (Scirpus cyperinus), has a definite ruderal character – its rhizomes are less substantial than those of Long’s bulrush (diameter: 1-1.5 cm vs. 1.5-3 cm, respectively), its leaves and culms are less robust, and its tussocks flower annually and do not spread aggressively (Whittemore and Schuyler 2003, Hill and Johansson 1992). According to Grime (1977), the wooly bulrush would be constitutively ruderal, i.e. completely adapted to disturbance. Wooly bulrush is common and widespread in eastern North America (Whittemore and Schuyler 2003) and Nova Scotia (Roland and Zinck 1998), occurring on the margins of wetlands and open still and running water, in forested swamp, and in ditches and other sites of anthropogenic disturbance, where soils are frequently but not necessarily continuously wet (personal observations). Long’s bulrush is an endangered species of the Atlantic Coastal Plain Flora (Wisheu et al. 1994). Its known range is from southwestern Nova Scotia to the pine barrens of southern New Jersey (Hill and Johansson 1992). Nova Scotian populations are of scientific interest – Has S. longii adapted to Nova Scotian conditions? What is the history of post-glacial migration of the species? – and of considerable concern in conservation. Long’s bulrush occurs in Canada only in southwestern Nova Scotia and may be genetically distinct from populations in the USA. Long’s bulrush is listed as vulnerable (Wisheu et al. 1994) under the Nova 3 Scotia Endangered Species Act (2006), of special concern under Canada’s Species at Risk Act (2006), and is globally imperiled (NatureServe 2010). Long’s bulrush is a High Priority species according to the Recovery Strategy and Management Plan for Multiple Species of Atlantic Coastal Plain Flora in Canada (Environment Canada and Parks Canada Agency 2010). This plan (p. 4) points out that, as ACPF populations in Nova Scotia are at the northern ends of their ranges and disjunct from populations in the USA, work is needed to determine the genetic distinctiveness and conservation-value of these populations. In 2010, MacKay et al. reported on the genetics of several populations of Long’s bulrush in the lower watershed of the Medway River in Queen’s County, Nova Scotia. One of these populations is unusual in that it includes patches (< 1% of the area covered by Scirpus) that flower annually (Hill 1994) and which show some morphological traits of the wooly bulrush. This suggested that Long’s bulrush and the wooly bulrush had hybridized to produce some plants that express the annual flowering-trait of the wooly bulrush putative parent species. MacKay et al. (2010) were able to show that these patches were indeed the product of hybridization. Unexpectedly, hybridization was also evident in the non-flowering part (> 99% of the area covered by Scirpus) of this population. Furthermore, hybridization was evident in one of two neighbouring (presumed “pure”) Long’s bulrush populations. Apparently, Long’s bulrush is vulnerable to genetic invasion by its sister species – S. longii and S. cyperinus are not yet reproductively incompatible despite being quite distinct in morphology, ecology and reproductive biology. Induction of flowering of Long’s bulrush (naturally or by human activity) puts this species at risk of loss by hybridization and introgression. Anthropogenic disturbance in the vicinity of Long’s bulrush populations may be especially threatening as disturbance induces flowering of the species and provides opportunity for the always-flowering wooly bulrush (MacKay et al. 2010). If reproductively compatible, only differences in flowering seasons (early summer for Long’s bulrush; late summer for the wooly bulrush; Roland and Zinck 1998) would inhibit the exchange of pollen between the species. The primary objective of this project is to determine whether an important population of Long’s bulrush in Nova Scotia has been compromised by hybridization with the wooly bulrush. A large population of Long’s bulrush is found in a wetland at the northern end of Lac de l’École in the main course of the Tusket River, Yarmouth County NS (Hill and Johansson 1992). The Tusket River watershed is a High Priority watershed in the Recovery Strategy and Management Plan for Multiple Species of Atlantic Coastal Plain Flora in Canada (Environment Canada and Parks Canada Agency 2010). Unusually, flowering occurs annually in the Lac de l’École population (Hill and Johansson 1992; personal observations in 2009 and 2010), suggesting (MacKay et al. 2010) that 4 (1) induction of facultative flowering of Long’s bulrush is occurring due to some regularly occurring environmental factors(s), and/or (2) at least some of the Scirpus in this population is the product of hybridization between Long’s bulrush and the wooly bulrush and annual flowering is caused by genetic input from the latter parent species. As the population is in an isolated area, it is unlikely that the annual flowering of Long’s bulrush at Lac de l’École, whether by (1) or (2), is due human activities. Thus, this population may be an example of natural hybridization – a “hybrid zone”. Demonstration of such hybridization would suggest Long’s bulrush has had a long history of exposure to genetic invasion by its sister species – a conclusion that would inform the conservation of Long’s bulrush, particularly efforts to isolate Long’s bulrush from its wooly relative. The technique developed for the investigation of the population genetics of Long’s bulrush in the Medway River watershed (MacKay et al. 2010) will be used to look for genetic markers of the wooly bulrush in flowering and non-flowering individuals of the Lac de l’École Long’s bulrush population. This simple technique involves isolation of small amounts of DNA from leaf tissue and characterization of that DNA by the RAPD (Random Amplified Polymorphic DNA) technique (MacKay et al. 2010). The work completed on the Medway River populations defined sets of species-specific genetic markers. This information will be combined with the Lac de l’École genetic information to determine whether some individuals at Lac de l’École are the products of hybridization. This project is divided into three practical parts: (a) surveying and sampling the Lac de l’École bulrush population, and sampling wooly bulrush in the vicinity of this population; (b) conducting genetic and morphological analyses; and (c) reporting the conclusions of these analyses to the bodies responsible for conservation of Long’s bulrush in Nova Scotia (The Atlantic Coastal Plain Flora Recovery Team and The Nova Scotia Nature Trust) and to the broader scientific community. These are described in full in NSSARCF09_05 (copy attached), with Project Objectives 1, 2 and 3 corresponding to (a) above, Project Objectives 4 and 5 corresponding to (b), and Project Objective 6 corresponding to (c). NSSARCF09_05 proposed a two-year project, with part (a) to be completed in summer 2010, part (b) to be completed by the end of 2011, and part (c) to be initiated in winter 2012. NSSARCF09_05 proposed that funding ($16,041.40) for this project would be divided 60% ($9,624.84) and 40% ($6,416.56) between the NSSARCF and Mount Saint Vincent University’s Internal Standard Research Grant program. The applicant was awarded $8,000 by the NSSARCF. At this time (May 2011), the recipient of NSSARCF09_05 has not applied for a related 5 MSVU internal grant, as his duties as departmental chair have restricted research activities in the past two years. The monies provided by the NSSARCF have been used to complete part (a) only. Only NSSARCF monies have been used for part (a). This report describes the work done to complete part (a). Plans for completing parts (b) and (c) are discussed in Summary and Prospects. 6 Logistics and Methods As proposed in NSSARCF09_05, three field trips were made to Lac de l’École in the summer of 2010. Lac de l’École is a small lake in the main course of the Tusket River, in Yarmouth County NS, immediately downstream from Gillifillan Lake and upstream from Wilson’s Lake. See W2 on the Tusket page of The Nova Scotia Atlas (1:150,000).
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